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APR 91992 

RIES NO. 42 


e Contribution of Archaeology 
to the Zoogeography of Borneo, 
with the First Record of a Wild Canid 
of Early Holocene Age 

Earl of Cranbrook 

A Contribution in Celebration 

of the Distinguished Scholarship of Robert F. Inger 

on the Occasion of His Sixty-Fifth Birthday 


March 31, 1988 
Publication 1385 


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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif, 943 pp. 
Grubb, P. J.. J. R. Lloyd, and T. D. Pennington. 1963. A comparison of montane and lowland rain forest 

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The Contribution of Archaeology 
to the Zoogeography of Borneo, 
with the First Record of a Wild Canid 
of Early Holocene Age 

Earl of Cranbrook 

Great Glemham House 
Suffolk. IP 17 1LP 

A Contribution in Celebration 

of the Distinguished Scholarship of Robert F. Inger 

on the Occasion of His Sixty-Fifth Birthday 

Accepted for publication March 31, 1986 
March 31, 1988 
Publication 1385 


© 1988 Field Museum of Natural History 

ISSN 0015-0754 


Table of Contents 

List of Tables 

Abstract 1 

Introduction 1 

Canid Remains from Agop Sarapad, Ma- 

dai Caves, Sabah 2 

The Specimens 3 

Discussion 4 

Identification of the Specimens 4 

Post-Pleistocene Extinctions in Borneo ... 5 

Acknowledgments 6 

Literature Cited 6 

Measurements of the left calcaneum of 
selected specimens of canid species 

List of Illustrations 

The dhole calcaneum and canine tooth 
from Agop Sarapad, Madai caves, Sa- 
bah 2 

The calcaneum positioned in calipers to 
measure maximum length 4 


The Contribution of Archaeology 
to the Zoogeography of Borneo, 
with the First Record of a Wild Canid 
of Early Holocene Age 


The extant fauna of Borneo lacks several large 
mammal species that are widespread elsewhere in 
the Sunda region. Archaeological research in Bor- 
neo has already found remains of three of these in 
late Upper Pleistocene or Holocene contexts: the 
tiger, Panthera tigris; Malay tapir, Tapirus indicus; 
and Javan rhinoceros, Rhinoceros sondaicus. Evi- 
dence of a fourth, the dhole, Cuon alpinus, is now 
reported from a midden dated about 10,000 b.p. 
in Agop Sarapad, Madai caves, Sabah. Extinction 
of these species evidently occurred within the last 
few thousand years and is attributed to failure to 
adapt to the environmental consequences of post- 
glacial climatic changes. 


In his monograph on the amphibians of Borneo, 
Robert Inger anticipated that new species would 
be discovered but that these additions would not 
materially affect conclusions on the geographical 
relations of the fauna (Inger, 1966, p. 357). Both 
predictions have since come true. Inger himself 
has subsequently described new Bornean taxa (In- 
ger & Frogner, 1979; Inger & Gritis, 1983), and 
Dring (1983a,b) has added others. Yet the overall 
picture of amphibian zoogeography in the region 
remains fundamentally unaltered. The fauna un- 
doubtedly originated from continental Asia and 
shows little affinity with Celebes, only a short dis- 
tance to the west, or with the Philippine Islands. 
There has, however, been a significant local ra- 
diation at lower taxonomic levels, recognizable in 

an assemblage of species confined to the Sundaic 
region; that is, the islands of the Sunda shelf and 
the Malay Peninsula south of about 10°N latitude. 
Species endemic to Borneo (comprising about 40% 
of the island's amphibian fauna) represent a spe- 
cial group among the Sundaic assemblage. Of the 
nonendemics, many species (about 80%) also oc- 
cur in Sumatra, somewhat fewer in Peninsular Ma- 
laysia (formerly Malaya), and only about 45% in 

Similar patterns characterize the zoogeography 
of the other terrestrial vertebrate classes. Thus, 
among the mammals, the fauna has clearly orig- 
inated from continental Southeast Asia, modified 
by a significant radiation within the Sunda region. 
Endemism is high in Borneo, with 37 (29%) of 1 29 
species of land mammals (i.e., excluding bats and 
marine mammals) currently being recognized as 
endemics. Of the 92 nonendemic species in Bor- 
neo, 87 (95%) also occur in Sumatra, 80 (87%) in 
Peninsular Malaysia, and only 44 (49%) in Java 
(data from Medway, 1977a, and Zon, 1979). 

Within the Sunda region, anomalous distribu- 
tions can be recognized, in several cases involving 
large mammal species which might be expected 
most easily to cross geographic barriers. In con- 
temporary Borneo, notable absentees are tiger, 
Panthera tigris, at present in Peninsular Malaysia 
(pm), Sumatra (s), and Java (j); leopard, Panthera 
pardus (pm, ?s, j); wild dog or dhole, Cuon alpinus 
(pm, s, j); Eurasian wild pig, Sus scrofa (pm, s); 
Javan rhinoceros, Rhinoceros sondaicus (pm, s, j); 
Malay tapir, Tapirus indicus (pm, s); and serow, 
Capricornis sumatraensis (pm, s). Among smaller 
mammals, only one giant squirrel of the genus 
Ratufa occurs (i.e., R. affinis), the widespread R. 
bicolor (pm, s, j) being absent. Also lacking are the 


bamboo rat, Rhizomys sumatrensis (pm, s), and 
the brush-tailed porcupine, Atherurus macrourus 
(pm, s). The banteng, Bos sondaicus, the only wild 
cattle in Borneo, is also native in Java, but is ab- 
sent from Sumatra and Peninsular Malaysia. Like 
the ferret badger, Melogale orientalis (j), B. son- 
daicus occurs in parts of mainland continental 
Southeast Asia, but only in Java and Borneo on 
the Sunda shelf. These two distributions, coupled 
with the occurrence in Borneo of a species of long- 
nosed ground squirrel, Dremomys, and a smooth- 
tailed treeshrew, Dendrogale (genera found else- 
where only on continental Southeast Asia), led 
Chasen (1940, pp. xi-xv) to propose a limited col- 
onization of the Sunda region by "eastern drift" 
from the Indochinese region. He also envisaged a 
"western drift from the continent by way of Su- 
matra" to explain the absence of the Eurasian wild 
pig, tiger, and dhole from Borneo. 

Archaeological evidence shows that neither pos- 
tulate is in fact required. It has long been known 
that the Pleistocene mammal fauna of Java in- 
cluded species now extinct on that island but sur- 
viving elsewhere in the Sunda region; for example, 
siamang, Hylobates syndactylus; orangutan, Pongo 
pygmaeus; Malay bear, Helarctos malayanus; el- 
ephant, Elephas maximus; bearded pig, Sus bar- 
batus; tapir; and bamboo rat. In Sumatra, banteng 
remains have been found in cave deposits attrib- 
uted either to a Middle Pleistocene interglacial or 
to the early Holocene (data reviewed by Hooijer, 
1975, and Vos, 1983). More recently, excavations 
by the Sarawak Museum in the 1950-1 960s and 
the Sabah Museum in the early 1980s have re- 
vealed the former presence in Borneo of four of 
the large mammals missing from the modern fau- 
na: (1) the tiger, represented at Niah cave, Sarawak 
(see Harrisson, 1972, and bibliography therein), 
by a deciduous canine from Neolithic levels 
(Hooijer, 1963); (2) the tapir, represented at Niah 
by a dozen pieces ranging in date from late Upper 
Pleistocene to about 8000 b.p. (Medway, 1960); 
(3) the Javan rhinoceros represented by a decid- 
uous molar and a fragment of ulna in a midden 
dated at about 10,000 b.p. (designated "MAD 2" 
by the excavator) in the Agop Sarapad mouth of 
Madai caves, Sabah (Bellwood, 1984), and (iden- 
tified with lessened confidence) by a left ectocu- 
neiform and a fragmentary lateral proximal pha- 
lanx from depths at Niah attributable to an early 
Holocene age (see Cranbrook, 1986, for details of 
these rhinoceros finds); and (4) the dhole, docu- 
mented below. The problem in these cases is no 
longer to elucidate routes or barriers to the colo- 

nization of Borneo, but rather to explain local ex- 
tinctions within the past few thousands of years. 

Canid Remains from Agop Sarapad, 
Madai Caves, Sabah 

The excavations were carried out in 1980 in the 
caves at Madai near Kunak, Sabah, 118°08'E, 
4°44'N, by staff of the Sabah Museum in associ- 
ation with Dr. Peter Bellwood (Bellwood, 1984, 
fig. 1; Bellwood, unpubl. data). Animal remains, 
including vertebrate teeth and bones, were sepa- 
rately bagged from each layer and later examined 
by myself in the museum premises at Kota Kin- 
abalu. Notes made at the time have been worked 
into a general report on animal remains, which 
has been placed on file in the museum. It is planned 
to publish an edited version as part of the exca- 
vator's final report (Bellwood, unpubl. data). Spec- 
imens appearing to be of particular interest (and 
therefore brought to England for further study) 
included the two described below. Both derive from 
the Agop Sarapad mouth (MAD 2), Madai caves, 
square HI, layer 2, at 5-10 cm. This provenance 
is firmly within the MAD 2 riverine shell midden, 
14 C-dated to about 10,000 b.p. (Bellwood, pers. 
comm.). The specimens are illustrated in Figure 
1 . Both have been returned to the Sabah Museum, 
Kota Kinabalu, Malaysia, for safekeeping. A cast 

Fig. 1 . The dhole calcaneum and canine tooth from 
Agop Sarapad, Madai caves, Sabah. Scale is in centi- 


Table 1 . Measurements of the left calcaneum of selected specimens of canid species. 

Measurementst (mm) 





Maximum Maximum 




reg. no. 






Archaeological specimen 

Agop Sarapad. Madai 







Malaysian pariah-type dog 









Cuon alpinus javanicus 








Cuon alpinus dukhunensis 








Canis familiaris dingo 








Canis familiaris dingo 








Canis lupus 








Canis aureus anthus 








* All examples are full-grown adults in the collection of the British Museum (Natural History). 

t Measurements were taken as follows: median length — from the tendinial groove on the superior face of the tuber 
calcanci to the inferior facet of the body; maximum length— the greatest length measured by calipers positioned as 
shown in Figure 2; maximum depth— the greatest distance between the anterior and posterior surfaces measured at 
the base of the tuber; maximum breadth— the breadth from the median edge of the sustentaculum tali to a point 
opposite it on the lateral surface on the body of the calcaneum; minimum breadth (tuber)— the mediolateral breadth 
of the tuber calcanei at its narrowest point. 

% A cast. The original has been returned to the Sabah Museum. 

of the calcaneum has been made by the British 
Museum (Natural History), where it is held in the 
Palaeontology Department under the registration 
number M 42838. 

The Specimens 

Right Lower Canine— Damaged, now con- 
sisting only of the root and dentine core of the 
crown. A very small area of enamel remains on 
the outer (labial) face at the base of the crown, 
shiny dark brown in color. This small piece of 
enamel is adequate to provide an orientation point 
from which the root can be measured, but is of no 
value for the specific identification of the tooth. 

Measurements are as follows: 

Root, length 23.3 mm 

Root, maximum breadth in anteroposterior 

(= mesiodistal) plane 9.7 mm 

Root, maximum breadth in lateral plane 

6.5 mm 

Left Calcaneum— Complete and undamaged. 
This bone is colored dark brown. It is indisputably 
attributable to a member of the dog family, Can- 
idae. Among likely species of Asian wild dogs, 
comparative measurements (table 1 ) show that this 
calcaneum is slightly larger in principal dimen- 
sions than its homologue in the skeleton of an 
adult male dhole of the Javan subspecies, Cuon 

alpinus javanicus, but smaller than that of an adult 
female of the Indian subspecies, C. a. dukhunensis. 
It is distinctly smaller than wolf, Canis lupus, and 
larger than jackal, Canis aureus (a member of the 
African race, C. aureus anthus, being the only 
available adult example). Neither is therefore in- 
dicated by the measurements. The discovery of 
either of these two Canis species in Borneo would 
represent a large extension of known range, his- 
toric or prehistoric. Both are thus best excluded 
from consideration. 

There remains the possibility that a type of do- 
mestic dog Canis "familiaris" is represented. 
Available for comparison is the skeleton of a fe- 
male of the primitive Malaysian pariah-type breed 
of domestic dog, an individual described and pic- 
tured by Medway (1977b, plate 3, 'Manggis'), now 
deposited at the British Museum (Natural Histo- 
ry); this is markedly smaller. Examples of Austra- 
lian dingo in the bm(nh) collections show wide 
variation in size; measurements of the calcanea of 
two adults bracket the archaeological specimen. 

Examination of a wider selection of calcanea of 
dholes and dingos does not reveal any nonmetric 
character that serves to distinguish the species. 
Features investigated included the relative dimen- 
sions of the tuber calcanei, the shape and angle of 
the sustentaculum tali, the angle of the ventral face 
of the process at the anterior base of the tuber, 
and the size and shape of the small facet at the 
mesioventral comer of the anterior face of the body 



Fig. 2. The calcaneum positioned in calipers to measure maximum length (table 1). 

of the calcaneum (terminology following Hughes 
& Dransfield, 1953). In no case was there any 
species-specific difference discernible in the sam- 
ples available. I conclude that, on morphology 
alone, the specimen could be attributed to a dhole 
or to a primitive, dingo-like form of domestic dog. 
Arguments for a more positive identification must 
be based on nonmorphometric grounds. 


Identification of the Specimens 

On the one hand, the dog was among the first 
of the mammals to be domesticated. In Europe 
and the Near East, remains attributed to dogs, 
rather than wolves, appear in association with Epi- 
paleolithic or Mesolithic cultures of early postgla- 
cial age. Dates suggested for the pioneer successes 
in domestication range from 12,000 b.p. (Davis & 
Valla, 1978) to 14,000 b.p. (Nobis, 1979). 

On the other hand, no such early dates have 
been established in Southeast Asia. In Borneo, re- 
mains of undoubtedly domesticated dogs have been 
found in archaeological contexts in other caves, 
but in none have cultural associations indicated 

an age earlier than Neolithic, at the oldest (Med- 
way, 1977b). In Thailand prehistoric sites have 
produced domestic dog bones dating back to 3500 
b.c. (Higham et al., 1980), a period considerably 
less ancient than the Agop Sarapad midden. 

Relevant evidence is the lack of other signs of 
the presence of domestic dog at these sites. While 
examining the bone from Agop Sarapad (and also 
the much larger quantity from deep levels at Niah), 
I have looked for the distinctive marks of gnawing 
by carnivores and found none. These middens 
originated as the food remains of human cave vis- 
itors. If contemporary man had been accompanied 
by domestic dogs, at least some of the bones must 
have been gnawed. I conclude that the most plau- 
sible identification for the two archaeological spec- 
imens is a wild dog, which itself was perhaps tl{e 
quarry of man. On the grounds of size and zoo- 
geography, the dhole is indicated. 

Intriguingly, as for tiger (Gersi, 1975) and tapir 
(Medway, 1977a), anecdotal accounts exist of the 
presence of the dhole in Borneo in historic times. 
Hose (1893, p. 26) included Borneo in the range 
of the species (under the name Cyon rutilans), 
commenting: "This wild dog must be very rare in 
Borneo. I have constantly heard native accounts 
of it, but I have never seen a specimen." Later, 
recounting a trip to Batu Bukit Song, Sarawak, 


Hose (1929, p. 144) reported sighting a "pair of 
wild dogs . . . they had been eating the remains of 
a young wild pig, but before we had time to get 
more than a glimpse of them they were away." At 
no time in the history of zoological collection has 
a specimen been taken. Hose's record was not ad- 
mitted by subsequent authors, including E. Banks 
and F. N. Chasen (see Chasen, 1940, p. 93), both 
of whom had much field experience in the region. 

Post-Pleistocene Extinctions in Borneo 

The complete list of 58 mammal species iden- 
tified in the excavations at Niah, including do- 
mestic dog and goat but excluding the Javan rhi- 
noceros (at that time unrecognized), has been 
published elsewhere (Medway, 1979, table 2). The 
total from Madai is smaller (Cranbrook, unpubl. 
ms. deposited at Sabah Museum, Kota Kinabalu). 
Both lists serve to confirm that the mammal fauna 
at the threshold of the Holocene era was very sim- 
ilar to that of present-day Borneo. 

The deepest levels at Niah yielded remains of 
one totally vanished mammal, the giant pangolin, 
Manis palaeojavanica (Hooijer, 1 960a), but this 
apparently did not survive into the Holocene era. 
The early Holocene fauna of Borneo presumably 
included the mammals already mentioned (tiger, 
dhole, tapir, and Javan rhinoceros) which have 
since become locally extinct. Over the same pe- 
riod, other species, which persist in modern Bor- 
neo, have undergone metrical changes. Thus, there 
has been a decline in tooth size among orangutans 
(Hooijer, 1960b) and monkeys of the genera Pres- 
bytis (including subgenus Trachypithecus) and 
Macaca (Hooijer, 1962). Among two species of 
giant rats, Rattus sabanus and R. muelleri, there 
have been changes both in the size and the relative 
dimensions of the teeth (Medway, 1964). Larger 
body size among the prehistoric populations is 
indicated by the dimensions of limb and foot bones 
of Sumatran rhinoceroses, Dicerorhinus suma- 
trensis (Cranbrook, 1 986), and barking deer, Mun- 
tiacus muntjak (Medway, 1959). 

The same archaeological studies show that man 
was present throughout the period concerned. Yet, 
as I have argued elsewhere (Medway, 1979), there 
are no grounds to suggest that direct or indirect 
impacts of human activity contributed to evolu- 
tionary changes or to the extinction of any species 
before the advent of the shotgun. Explanations 
must be sought in natural ecological processes. 

Advances in palaeogeographical research have 

improved our understanding of events in the re- 
gion during the Cenozoic and subsequently. The 
antiquity of the Makassar Strait is confirmed, al- 
though it is now recognized that Celebes has a 
complex tectonic origin (Audley-Charles, 1981). 
During the Pleistocene there were large extensions 
in the exposure of subaerial land of the Sunda shelf 
associated with glacial episodes, as summarized 
by Inger (1966) from earlier sources. It is also 
known that these were accompanied by climatic 
changes sufficient, for instance, to cap Mt. Kina- 
balu with permanent ice (Koopmans & StaufTer, 
1967). It seems that present-day conditions in the 
Sundaic region are atypical of the Quaternary as 
a whole, with sea level and the upper forest limit 
on mountains exceptionally high, average tem- 
peratures close to maximum Quaternary values, 
and the climate at or near its wettest (Whitmore, 
1 98 1 ). A change from cooler, drier, and more sea- 
sonal conditions apparently took place rather rap- 
idly in the first millenia of the Holocene, accom- 
panied by rising sea levels which reached a 
maximum of at least 6 m above present shorelines 
about 5000 b.p. (Haile, 1971). 

The cooler and more seasonal climate of Pleis- 
tocene glacial periods, including the last, which 
immediately preceded the Holocene intermission, 
is presumed to have favored the development of 
a mosaic of forest broken by open gaps and glades. 
Under such conditions, the forest-edge facies must 
have been extensive, with abundant browse ac- 
cessible to ground-dwelling ungulates. Following 
the change to a warmer and wetter postglacial cli- 
mate, the open spaces would have been invaded 
by closed forest. Mature \ropical evergreen rain 
forest supports a sparse ground and shrub storey 
vegetation. In the deep shade, plant life is restrict- 
ed. The main zone of productivity is in the upper 
canopy, high out of reach of ground-dwelling her- 
bivores. These changes would therefore have de- 
graded the quality of the environment for all non- 
scansorial phytophagous mammals of the ground 
or shrub storeys, and for large-bodied browsing or 
grazing perissodactyls in particular. Indirect con- 
sequences would have impinged upon nonscan- 
sorial predators of these mammals, again espe- 
cially the large-bodied carnivores. The affected 
populations would thus have experienced ecolog- 
ical pressures simultaneously with the final sev- 
erance of land connections across the Sunda shelf 
and hence isolation from sources of new genetic 

Nevertheless, Bornean populations of other 
mammal species have continued from the early 


postglacial to the present. Among these, the beard- 
ed pig, Sus barbatus, as far as archaeological ma- 
terial can show, has undergone no morphological 
change (Medway, 1978). At the same time, the 
Sumatran rhinoceros and barking-deer have re- 
sponded to the changing environment by reduc- 
tion in individual body size, a modification rec- 
ognized as adaptative to restricted resources. The 
metrical changes in the teeth of other mammals, 
as noted above, involving absolute and/or relative 
dimensions, may also reflect alterations in body 

The archaeological evidence is inadequate to 
demonstrate changes in abundance. Populations 
affected by postglacial alterations in the environ- 
ment were likely to have declined in response to 
the diminution of accessible resources. Large 
mammals might already have been comparatively 
rare and hence vulnerable to local extinction, even 
on so big an island as Borneo. Reduced population 
density among the primary consumers has inevi- 
table repercussions on members of the community 
at higher trophic levels. It is therefore not unex- 
pected that the mammal species to become extinct 
in Borneo during the postglacial period included 
these four obligatory ground-dwellers: two large 
browsing ungulates (Javan rhinoceros and tapir) 
and two large predatory carnivores (tiger and 

I would be venturing too far from my own ground 
if I were to speculate on the likely effects of these 
environmental changes on amphibian faunas of 
the terminal Upper Pleistocene of Borneo. This 
train of thought I hereby offer to Bob Inger, with 
respect and affection, leaving him to pursue it as 
far as may be profitable in the light of his own 
studies of the existing fauna. 


I am grateful to Dr. Peter Bellwood for supplying 
radiometric dates, to Dr. Juliet Jewell for provid- 
ing facilities for this study at the British Museum 
(Natural History), and to both, with Dr. D. A. 
Hooijer, for kindly reading and commenting on 
this text in progressive stages of drafting. 

Literature Cited 

Audley-Charles, M. G. 1981. Geological history of 
the region of Wallace's line, chap. 4. In Whitmore, T. 

C, ed., Wallace's Line and Plate Tectonics. Clarendon 
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Bellwood, P. 1984. Archaeological research in the 
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Pacific Prehistory Association, 5: 38-56. 

Chasen, F. N. 1940. A handlist of Malaysian mam- 
mals. Bulletin of the Raffles Museum, Singapore, 15: 
i-xx, 1-209. 

Cranbrook, Earl of. 1986. A review of fossil and 
prehistoric remains of rhinoceroses of Borneo. Sabah 
Museum and Archives Journal, 1(1): 50-1 10. 

Davis, S. J. M., and F. R. Valla. 1978. Evidence for 

u the domestication of the dog 12,000 years ago in the 
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Dring, J. 1983a. Frogs of the genus Leptobrachella 
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. 1983b. Some new frogs from Sarawak. Am- 
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Gersi, D. 1975. Dans le jungle de Borneo (Kaliman- 
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Haile, N. S. 1971. Quaternary shorelines in West Ma- 
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Harrisson, T. 1972. The prehistory of Borneo. Asian 
Perspectives, 13: 17-45. 


1980. An analysis of prehistoric canid remains from 
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Hoouer, D. A. 1 960a. The giant extinct pangolin {Manis 
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. 1960b. The orang-utan in Niah cave prehis- 
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1 962. Prehistoric bone: The gibbons and mon- 

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. 1963. Further "Hell" mammals from Niah. 

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Inger, R. F., and P. A. Gritis. 1983. Variation in 
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1 977b. The ancient domestic dogs of Malaysia. 

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. 1979. The Niah excavations and an assessment 

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